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World Catalog of the Genera of Pselaphidae (Coleoptera)

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World Catalog of the Genera of Pselaphidae (Coleoptera) World Catalog of the Genera of Pselaphidae (Coleoptera) Abstract All names that have been proposed above the resolution of conflicting regional classifications, are species level in the family Pselaphidae are cata­ discussed, but only one new change is made; Aux- loged. with a full bibliography of the original de­ enocerini Jeannel is synonymized under Jubini scriptions and a full taxonomic index. The ap­ Rafiray. Application of the principle of priority to proximate world distribution and number of family-group names requires the following changes species of each currently recognized genus are giv­ in recent usage: Bythininae to Goniacerinae, Fus- en. along with the present status, type species, and tigerini to Clavigerodini, Octomicrini to Dimerini. mode of type species designation for all genus- Pyxidicerini to Bythinoplectini, and Zethopsina to group names. Bythinoplectina. Citations for family-group names Changes and problems at the generic level are are given in a separate list which serves as a con­ summarized in a separate Discussion section. New spectus of the suprageneric classification. names are proposed for 18 newly discovered ju­ Since the last world catalog of Recent Psela­ nior homonyms representing currently recognized phidae (Raffray, 191 1 ), the family has grown from pselaphid genera; six additional homonyms have 3,300 species in 468 genera to about 8,377 species available names. Four unavailable names in cur­ in 1,091 genera, placed in two divisions, six rent use are validated by designation of a type subfamilies, 10 sections, 57 tribes, and 40 sub­ species for each; 13 additional unavailable names, tribes. An additional 16 genera and 36 species of all of Karaman and most recently cited as syn­ extinct taxa, and two genera with two species based onyms, are listed but not validated. Type species on larvae, are listed. are designated for the first time for 21 older names. Previously unnoticed type species designations make Fustigerodes a senior objective synonym of Commatoceropsis, and Ramecia a senior objective synonym of LinioHs; Xovoclaviger is available for Introduction Fustigerodes of recent usage, and a new genus Ra- cemia is proposed forRamecia of recent usage. The family Pselaphidae, with about 8,400 de­ Rhvtus Westwood (not preoccupied) replacesAr- scribed species, is among the ten largest beetle hytodes Reitter. Problems with previous type families, but it is easily the least well known of species designations for Datniodes, Dabnoplectus, these families. This is undoubtedly due in large and Fustiger are reviewed. Unresolved problems measure to the small size of the beetles (usually 1 involving the forgotten name Alytus, previously to 3 mm) and their reclusive habits. Despite a overlooked type species designations forBatrisus, general scarcity of specimens in most insect col­ and the identity of the type species of Rybaxis are lections. the group has attracted the attention of discussed. A list of genera recently transferred to several very productive taxonomists, perhaps due or from Pselaphidae is also given. to the intriguing variation in external structure The genera are placed into higher taxa according which often includes striking cuticular modifica­ to the authors’ estimate of a currently accepted tions of the males. Use of modern collecting meth­ world classification, based on that of Jeannel. ods (Berlese/Tullgren funnels, Winkler/Moczarski Problems with this classification, including the extractors, blacklight and flight-intercept traps) is NEWTON & CHANDLER: CATALOG OF PSELAPHIDAE now providing a wealth of material, and interest family Staphylinidae (32,000+ named species), in the family is higher than ever among system- most members of which resemble pselaphids in atists as well as other biologists. An understanding having short elytra. Pselaphids differ from most of the present state of knowledge of the classifi­ staphylinids in having a reduced tarsal formula cation and distribution of the family is greatly (all three- or two-segmented), somewhat clubbed hampered, however, by the lack of any worldwide antennae, and a distinctive body form with a rel­ summary since the last catalog of the family (Raf- atively rigid abdomen. They also have a unique fray, 1911). The present work attempts to provide system of distinctive cuticular foveae, highly mod­ such a summary above the species level. ified female genitalia, and other adult and larval Knowledge of the natural history of pselaphids characters which imply that the family is mono- is usually, at best, restricted to the habitat in which phyletic. Pselaphids are generally considered to be the adults are collected. Most species are found in derived from staphylinids, with the main source leaf and wood litter of the forest floor (e.g., Park, of argument being to which subgroup of Staphy­ 1947a; Tanokuchi & Itoh, 1983; Chandler, 1987a), linidae they are most closely related. It is likely usually in low numbers of individuals. Species that, when a more phylogenetically defensible richness in this habitat may be very high, however, family classification of Staphylinoidea is adopted, with 30 or more species often turning up in litter Pselaphidae will be combined with at least part of samples from small plots of warm temperate and the present family Staphylinidae, as already done tropical forests (pers. obs.). Species can also be by Naomi (1985). For general discussions on the discovered in wetland, grassland, desert, beach, relationship between these two families and the cave, and even arboreal habitats, wherever debris, difficulties of distinguishing them, see Casey ( 1894), moss, or root mats maintain a zone of high hu­ Raffray (1908a), Scheerpeltz (1926), Park (1942), midity. A number of species scattered through five Jeannel (1950), Kasule (1966), Naomi (1985), and of the subfamilies are inquilines in the nests of Thayer (1987). The anatomy of the Pselaphidae social insects, while all species of the sixth subfam­ is discussed by Raffray (1908a), Park (1942), and ily, the Clavigerinae, are thought to be obligate Jeannel (1950). associates of ants or termites (Kistner, 1982). There have been two periods of explosive growth No complete life histories have yet been worked in the number of described species, genera, and out. The immature stages are very poorly known, higher taxa of Pselaphidae. The first period ex­ with larvae of fewer than a dozen genera and pupae tended from the late 1870s to about 1918, and was of three genera described (see especially Kemner, largely the result of the work of three men: Achille 1927; Böving & Craighead, 1931; Besuchet, 1952, Raffray, Edmund Reitter, and Ludwig Schaufuss. 1956b; Kasule, 1966; De Marzo & Vit, 1982; De All three produced numerous descriptive taxo­ Marzo, 1986; Newton, in press). Both larvae and nomic papers dealing primarily with faunal col­ adults are assumed to be predatory on small ar­ lections from various parts of the world, and each thropods and worms, based on limited observa­ also produced one or more attempts at proposing tions (Park, 1947a; Engelmann, 1956; Reichle, a higher classification of the family. Two large 1969; De Marzo, 1986) and on general mouthpart synthetic works by Raffray on the world fauna of structure and gut contents. the family, in the Genera Insectorum (1908a) and Members of the family are of no known com­ Coleopterorum Catalogus (1911) series, summa­ mercial importance, but the high species richness rize the results of this first active period. The fam­ and abundance in certain microhabitats suggests ily grew from 450 species in 44 genera (Gemmin- that pselaphids may be ecologically important ger & Harold, 1868) prior to this period, to 3,300 (Park, 1947a; Chandler, 1987a). This diversity, in species in 468 genera and 19 tribes (Raffray, 1911). combination with a wealth of taxonomic charac­ The second very active period ran from about ters that should facilitate phylogenetic analysis, 1942 to 1964 and was dominated by two men: also suggests a high potential for use of pselaphids Orlando Park and the remarkably prolific René in various evolutionary studies such as local and Jeannel. This period is also characterized primar­ large-scale biogeography and integration of guests ily by the descriptive taxonomy of faunas, with into social insect systems. each author confining his work to one faunal re­ Pselaphids have been maintained as a distinct gion for long periods of time. Park worked initially family of stable limits since the early part of the on the Neotropical fauna, and in his later years on 19th century. The family is placed in the super­ the Nearctic and portions of the Pacific faunas. family Staphylinoidea, adjacent to the even larger Jeannel initially worked on the western European 2 FIELDIANA: ZOOLOGY fauna, following with extensive work on all por­ Methods tions of the African fauna. Later in his career he touched on the Japanese, Chilean, and to a limited This catalog was undertaken in part to deter­ extent the Oriental faunas. mine the feasibility of using a microcomputer da­ The major workers following this second period tabase for constructing generic checklists for the have generally continued a regional approach but families of Staphylinoidea more easily than could have increasingly emphasized thorough revision­ be done on paper or with a word processor. The ary studies of more restricted taxonomic groups. basic procedure was to (1) set up a suitable data­ These include C. Besuchet (Palearctic), N. Leleup base (a commercially available database program, (African), I. Löbl (Oriental), and R. O. Schuster dBASE II®, was used on Morrow® and Norths- and A. A. Grigarick (Nearctic). G. Coulon (1986) tar® CP/M® microcomputers); (2) begin with the has completed the first worldwide revision of the last world catalog of the family (Raffray, 1911), genera and higher taxa of a sizeable group (Faron- entering all generic names and relevant data in­ inae excluding Faronini). Grigarick and Schuster cluding distribution and number of species for each (1980) and Coulon (1986) have presented the first recognized genus; and (3) go through the Zoolog­ explicit phylogenetic analyses of certain groups in ical Record in sequence from 1911 to 1986, adding the family.
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